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Revista de Biología Tropical

versión On-line ISSN 0034-7744

Rev. biol. trop vol.60  supl.3 San José nov. 2012

 

Marine biodiversity of an Eastern Tropical Pacific oceanic island, Isla del Coco, Costa Rica

Jorge Cortés1*,2*

*Dirección para correspondencia:


Abstract

Isla del Coco (also known as Cocos Island) is an oceanic island in the Eastern Tropical Pacific; it is part of the largest national park of Costa Rica and a UNESCO World Heritage Site. The island has been visited since the 16th Century due to its abundance of freshwater and wood. Marine biodiversity studies of the island started in the late 19th Century, with an intense period of research in the 1930’s, and again from the mid 1990’s to the present. The information is scattered and, in some cases, in old publications that are difficult to access. Here I have compiled published records of the marine organisms of the island. At least 1688 species are recorded, with the gastropods (383 species), bony fishes (354 spp.) and crustaceans (at least 263 spp.) being the most species-rich groups; 45 species are endemic to Isla del Coco National Park (2.7% of the total). The number of species per kilometer of coastline and by square kilometer of seabed shallower than 200m deep are the highest recorded in the Eastern Tropical Pacific. Although the marine biodiversity of Isla del Coco is relatively well known, there are regions that need more exploration, for example, the south side, the pelagic environments, and deeper waters. Also, several groups of organisms, such as the flatworms, nematodes, nemerteans, and gelatinous zooplankton, have been observed around the Island but have been poorly studied or not at all.

Key words: Marine biodiversity, Costa Rica, Isla del Coco, Cocos Island, Eastern Pacific, endemic species.

Resumen

La Isla del Coco es una isla oceánica en el Pacífico Tropical Oriental; es parte del Parque Nacional más grande de Costa Rica y es un sitio de Patrimonio Mundial. La isla ha sido visitada desde el Siglo XVI por su abundancia de agua dulce y árboles. Estudios de biodiversidad marina de la isla empezaron a finales del Siglo XIX, con un intenso período de investigación en la década de 1930, y de nuevo desde mediados de la década de 1990 al presente. La información sobre organismos marinos se encuentra dispersa y en algunos casos en publicaciones antiguas. En el presente trabajo se recopilan todos los registros publicados de organismos marinos de la  isla. Al menos 1688 especies han sido registradas, con los gasterópodos (383 especies), peces óseos (354 spp.) y crustáceos (al menos 263 spp.) como los grupos con más especies; de esas, 45 son especies endémicas del Parque Nacional Isla del Coco (2.7% del total). El número de especies por kilómetro de costa y por kilómetro cuadrado de lecho marino de menos de 200m de profundidad son los más altos de cualquier sitio estudiado. Aunque se conoce relativamente bien la  biodiversidad marina de la Isla del Coco,  hay regiones, por ejemplo, el lado sur, los  ambientes pelágicos, y las zonas más  profundas que requieren de más exploración. También, varios grupos de organismos han sido observados en la isla pero muy poco estudiados o no del todo, por ejemplo los gusanos planos, nemátodos y el plancton gelatinosos.

Palabras clave: Biodiversidad marina, Costa Rica, Isla del Coco, Pacífico Oriental, especies endémicas.

Isla del Coco (Cocos Island) is an oceanic island in the Eastern Tropical Pacific Ocean, about 500km from mainland Costa Rica (Cortés 2008), and biogeographically it is part of the  Ocean  Island  Province  (sensu  Robertson & Cramer 2009). It is a highly diverse area with the highest number of endemic species in Costa Rica (Cortés & Wehrtmann 2009, Cortés 2013a). It was declared a National Park in 1978, a UNESCO World Heritage Site in 1997, and a Ramsar site in 1998. It is important for Costa Rica, due to of its biological and historical richness and also because the Territorial Sea surrounding the island makes the marine area of Costa Rica more than 10 times its terrestrial area (Cortés & Wehrtmann 2009, Cortés 2013b).

Marine  biodiversity  studies  at  Isla  del Coco started in the late 19th Century with the expedition of the US Fisheries Commission Steamer Albatross (see a history of marine research at what is now Parque Nacional Isla del Coco (Isla del Coco National Park) in Cortés 2008). Since then, many studies have been done but there is still more to be discovered. Hertlein (1963) compiled what was known about marine species of the Island. In this paper I update the list of marine organisms of Isla del Coco National Park, compare it with the species from the mainland, and identify areas for future research.

Materials and Methods

Published   records   of   marine   species of  Parque  Nacional  Isla  del  Coco  (PNIC) were searched and species lists compiled. An important source was the book “Marine Biodiversity of Costa Rica, Central America” edited by Wehrtmann and Cortés (2009). Here I included overlooked and newly published records. The number of species from PNIC is compared to the numbers from the rest of the Pacific of Costa Rica. Including the mainland, the Coco Volcanic Cordillera, the pelagic areas, and the Costa Rica Thermal Dome.

The perimeter of Isla del Coco is 28.8km, and it is surrounded by a platform that drops off at around 180-200m. The perimeter at 100m is 54.4km and at 200m is 71.4km. The area less than 50m deep around the island is 91.5km2, less than 100m is 133km2, and less than 200m deep  is  318km2  (O.G.  Lizano,  pers.  comm. 2011).  The  Linear  Biodiversity  Index  (LBI) and the Area Biodiversity Index (ABI) were calculated as in Wehrtmann et al. (2009). To calculate the LBI, the number of species was divided by the perimeter of the island. The ABI was calculated by dividing the number of species by the area down to the 200m isobaths; the depth within which almost all the species had been recorded.

Marine Biodiversity of Isla del Coco National Park

At least 1688 species of marine organisms (Appendix 1) have been reported from Isla del Coco National Park (PNIC). The most speciesrich groups are the gastropods (383 species), bony fishes (354 spp.) and crustaceans (at least 263 spp.) (Table 1). Close to 4,700 species of marine organisms have been reported for Costa Rica (Table 1); of these species, 747 or ~16% have been only reported from PNIC but not from other areas of Costa Rica. The percentages by taxonomic group ranged from 0 to 100%. All brown algae, echiuran, marine insects, and reptiles from PNIC are also found in the rest of Costa Rica. While all reported branchiopods, ostracods, phoronids, and crinoids, represented by one or two species, known from Costa Rica are reported from PNIC. Some groups have a disproportionate percentage of known species at PNIC, scyphozoans (40%), echinoderms as a phylum (42.6%), and its classes, asteroids (66.7%) echinoids (36.4%) and holothurians (40.4%), brachiopods (75.0%), and within the chordates, cephalochordates (50%) appendicularians (70%) and thaliacians (75%).

Species of several groups have been reported from other areas of Costa Rica than PNIC (Table 2). Some of these taxa we know are absent from PNIC, for example the seagrasses and mangroves, and some of the species associated to this ecosystems. Other groups have been observed, photographed or collected but there are no published accounts of them. Within these groups we have nematodes, nemerteans, ascideans, and parasites of fishes and turtles. Free-living flat worms have been observed along the mainland coast of Costa Rica as well as at PNIC, but there are no publications. Of other taxa we do not know if they are present or not, for example, marine fungi, cumaceans and kinorhynchans (Table 2). Plus there must be other marine groups that have been reported from the Eastern Tropical Pacific, for example, loriciferans (Heiner & Neuhaus 2007) that might be present at PNIC.

Forty five species or 2.7% of the species known  from  PNIC  are  endemic  (Tables  3, 4), and this represents 47.4% of all endemic marine species of Costa Rica (95 spp.). The number of endemic species is relatively low, but that is common in marine environments. The list of endemic marine species is presented in Table 3, as well as the reference to the publication where the species was described. Between 1893 and 1971, 16 species were described, while 29 were described from 1981 to 2011. Most endemic species are fishes (33.3% of all endemics from PNIC) and most were described in the last 30 years (11 of the 15 species). Crustacea is the next group with most endemism, 28.9%, followed by the mollusks, 15.5%, all very well studied groups (Table 4). Within a particular group, the brachiopods have the highest percentage of endemism, 16.7% followed by the sponges and polyplacophorans with 12.5% (Table 4).

Biodiversity indices used to compare species diversity at Isla del Coco with that at the Costa Rican coast revealed significantly higher values at PNIC than at the coast. For example, the Linear Biodiversity Index for Isla del Coco is 58.6, which is significantly higher than the highest  value  (3.8)  found  by  Wehrtmann  et al. (2009) for the Costa Rican Pacific coast. The same happens when comparing the Area Biodiversity Index of continental shelves. The value for the 200m isobaths of PNIC is 5.3, compared to the highest value of 0.3 reported by Wehrtmann et al. (2009). The LBI and ABI values for the Costa Rican coastline and continental platform, respectively, were the highest when compared to other countries in the region (Wehrtmann  et  al.  2009).  By  all  measures Isla del Coco is a very rich area in the eastern tropical Pacific.

Status of the endemic species

Forty five species are endemic to PNIC (Table 2). While some are abundant, such as the calcified hydrozoan Stylaster cocosensis, described in 1991 by Stephen D. Cairns, others have not been seen since they were described, for example, the sand dollar, Encope cocosi. This species had not been found alive since it was described by H.L. Clark in 1948. However,  in  January  2007  a  specimen  recently dead was dredged from deep water. In 1986 using the research submersible Johnson-SeaLink (Harbor Branch Oceanographic Institution,  Fort  Pierce,  Florida,  USA)  diving  to several hundred meters, collected specimens that resulted in new species and some were endemic (Cairns 1991a, b). We don not know the status of some of those endemics because no submersible with the depth capacity of the Johnson-Sea-Link has been back to the island. There  is  now  another  submersible  operating more regularly at the island, the DeepSee (Undersea Hunter Group, Puntarenas, Costa Rica), with a depth capability of 450m (Cortés & Blum 2008). We have been able to observed several  of  the  deepwater  endemics  collected in 1986, some are relatively abundant. Eleven endemic species have been described in the last decade so it’s possible that eventually they will be found in other areas. Some species of fishes which were initially classified as endemic to one of the oceanic islands of the eastern Tropical Pacific are now reported from one or more of the other oceanic islands. For example, Stegastes arcifrons which have been found in the three oceanic islands, Galápagos, Malpelo and Isla del Coco or Serranus tico and Halichoeres discolor, found in Isla del Coco and Malpelo. Starr et al. (2012) indicated in their study of deepwater fishes of Isla del Coco National Park and Las Gemelas Seamount that probably deep areas in the eastern tropical Pacific will have similar species, but more studies and collections are needed.

Discussion

Isla del Coco National Park has a rich marine biodiversity with some groups having been studied for many years and numerous scientists. For example, fishes and mollusks, especially gastropods, are relatively well known while other groups such as cyanobacteria, gelatinous zooplankton, nematodes and flatworms  have  been  poorly  studied  or  never at all even though we know they are on the island. As a result of recent expeditions (2006-2012) many new records of species have been reported (Dean et al. 2010a, 2012, Sibaja-Cordero et al. 2012), including a phylum, Phoronida (Dean et al. 2010b), and new species are being discovered, even of conspicuous groups such as octocorals (Breedy & Cortés 2011, Breedy et al. 2012). Reports of new records and descriptions of new species are being prepared at the present time.

Hertlein (1963) did a compilation of published marine species of Isla del Coco, and included a biogeographic analysis of the flora and fauna of the island, plus an annotated bibliography. He reported 334 species (Table 5), with the gastropods (62 species) as the most species-rich group, followed by bony fishes (59) and crustaceans (56). The number of species and of different taxonomic groups has increased  significantly  but  the  same  pattern of the most species-rich groups is maintained. Wehrtmann et al. (2009) reported 1,142 marine species for Isla del Coco National Park, with the most species-rich groups, in the same order, being the same as above. Here, 546 more species were added to the list of marine species of Isla del Coco National Park, and more will be added in the near future as other groups, depths and areas of the island are being studied.

Hickman (2009), in his study of the marine invertebrate  biota  of  the  Galápagos  Islands, found that while some groups of species are depauperated others displayed high diversity when compared to mainland Ecuador. Similar patters were observed at Isla del Coco National Park. These patterns can be attributed to several possible factors likely acting in concert, both for source populations from elsewhere as well as established populations at PNIC: variation in the dispersal potential to and from PNIC, the probability of recruitment at PNIC, and the potential for survival and continued recruitment based on local environmental conditions. Species with long-lived larvae will have a chance of dispersing more than others if they find the type of environmental conditions necessary to survive and reproduce. For example, the absence of seagrasses and the low number of species of bivalves may be due to the lack of soft sediments where they can live.

Areas for future research

The least studied area of PNIC is the south side due to the normally rough sea conditions on that side (Lizano 2008). From a few observations, several species and environments in the south are different from the north in species density and composition, probably due to the currents that flow there (Cortés & Blum 2008). More sampling should be done on that side in the future in the shallow and deepwaters of the south for better understanding the biodiversity of PNIC, and the effect of currents on that biodiversity.

There are several groups of organisms which have been observed and in some cases collected but for which there are no publications. Examples include cyanobacteria, sponges, flatworms, and nematodes (Table 2). For a few groups, especially the best known, there are some publications on their biogeographic relationships. Several species of stomatopods (Manning 1972), most reef building corals (Cortés 1986, 2011, Glynn & Ault 2000), some mollusks (Montoya & Kaiser 1988), sea urchins (Lessios et al. 1998), and about one third of the shore-fishes (Robertson et al. 2004) are related western Pacific species. More molecular work is needed to discover cryptic species (e.g. Knowlton 2000, Boulay et al. in prep.), and the genetic connectivity (e.g. Lessios & Robertson 2006) of PNIC populations with other areas.

Polidoro et al. (2012) indicated the importance of species-specific information regarding population trends and extinction risks for developing conservation strategies. To do this we must first know what is there, which this paper intends to fulfill. Then we need to know what is the status of the populations, how they are changing over time, and what is affecting them. Unfortunately for most groups this information is unknown.

Acknowledgements

I thank the following scientists for their help with their group of specialty and/or review of  the  manuscript:  Fabián Acuña,  Juan  José Alvarado, Peter Auster, Gilbert Barrantes, Odalisca Breedy, Richard Brusca, William Bussing, Allan Carillo, Allen Collins, Harlan Dean, Ana Dittel, Cindy Fernández, Cristian Pacheco, Christian Emig, José Leal, Laurence Madin,  Ross  Robertson,  Eva  Salas,  Astrid Sánchez, Jeffrey Sibaja-Cordero, Rick Starr, Robert  van  Syos,  Benjamin Victor  and  Rita Vargas. Omar Lizano for providing the data on perimeter and areas of the marine sections of Isla del Coco National Park (PNIC). Research at PNIC has been funded by the Vicerrectoría de Investigación and CIMAR of the Universidad de Costa Rica, Conservation International (CI), Fonds Français pour l’Environnement Mondial  (FFEM)  and  the  Consejo  Nacional de Rectores de las Universidades Públicas de Costa Rica (CONARE). The preparation of this publication was advanced significantly during my stay as visiting Professor at Newcastle University, Newcastle, United Kingdom. Support has been received from Área de Conservación Marina Isla del Coco (ACMIC) and the Undersea Hunter Group.

References

1. Abele, L.G. & W. Kim. 1984. Notes on the freshwater shrimps of Isla del Coco with description of Macrobrachium  cocoense,  new species.  Proc.  Biol.  Soc. Washington 97: 951-960.         [ Links ]

2. Acuña, F.H., J. Cortés & A. Garese. 2012. Ocurrence of the sea anemone Telmatactis panamensis (Verrill, 1869.  (Cnidaria:  Anthozoa:  Actiniaria.   at   Cocos Island National Park, Costa Rica. Rev. Biol. Trop. 60 (Suppl. 3): 201-205.         [ Links ]

3.  Allen, G.R. & D.R. Robertson. 1994.  Fishes of the Tropical Eastern Pacific. Univ. Hawaii Press, Honolulu, Hawai’i. 332 p.         [ Links ]

4. Allen, G.R. & D.R. Robertson. 2002.  Halichoeres salmofasciatus, a new  species  of wrasse (Labridae) from Isla del Coco, tropical eastern Pacific. Aqua 5: 65-72.         [ Links ]

5. Alvarado, J.J. 2010. Isla del Coco (Costa Rica) Echinoderms: State of  knowledge, p. 103-113. In: L.G. Harris,  S.A.  Böttger,  C.W. Walker  &  M.P.  Lesser (eds.). Echinoderms: Durham.  Proc. 12th Int. Echinoderm Conf. CRC Press, Taylor & Francis Group, Balkema, Leiden, Netherlands.         [ Links ]

6. Alvarado, J.J. & A. Chiriboga. 2008. Distribución y composición de los equinodermos de las aguas someras en la Isla del Coco, Costa Rica (Pacífico Oriental). Rev. Biol. Trop. 56 (Suppl. 2): 99-111.         [ Links ]

7. Alvarado, J.J. & J. Cortés. 2009. Echinoderms. Text: 421-433,  Species list, CD: 392-408. In:  I.S. Wehrtmann  &  J.  Cortés  (Eds.).  Marine  Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

8. Bakus, G.J. 1975. Marine zonation and  ecology of Cocos Island, off Central America. Atoll Res. Bull.179: 1-11.         [ Links ]

9. Bamber, R.N. 2009. Sea-spiders. Text: 307-311, Species list, CD: 237-238. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine  Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

10. Barham, E.G. 1979. Giant larvacean houses: observations from deep submersibles. Science 205: 1129-1131.         [ Links ]

11.Barnard, J.L. 1991. Amphipoda of the  Galápagos Islands. Pp. 193-206. In: M.J. James (ed.), Galápagos Marine Invertebrates: Taxonomy, Biogeography, and Evolution in Darwin´s Islands.  Plenum Press, New York and London.         [ Links ]

12. Barrantes, G. & J. Chaves-Campos.  2009. Birds in coastal and marine environments. Text 469-478, Species list, CD: 479-484. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of  Costa Rica, Central America. Springer, Berlin.         [ Links ]

13. Bernecker, A. 2009. Marine benthic algae. Text 109-117, Species list, CD: 17-70. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

14. Bernecker, A. & I.S. Wehrtmann. 2009. New records of benthic marine algae and Cyanobacteria for Costa Rica, and a comparison with other Central American countries. Helgol. Mar. Res. 63: 219-229.         [ Links ]

15. Bigelow, H.B. 1909. Reports of the  scientific results of the Expedition to the  Eastern Tropical Pacific, in charge of Alexander Agassiz, by the U. S. Fish Commission  Steamer  “Albatross,”  from  October,  1904, to  March,  1905,  Lieut.  Commander  L.  M.  Garrett, U.S.N., commanding. XVI. The Medusae. Mem. Mus. Comp. Zoöl., Harvard Coll. XXXVII: 1-243.         [ Links ]

16. Bigelow, H.B. 1928. Scyphomedusae from the Arcturus Oceanographic Expedition. Zoologica 8: 495-524.         [ Links ]

17. Bigelow, H.B. 1931. Siphonophorae  from the Arcturus   Oceanographic   Expedition.   Zoologica   8: 525-592.         [ Links ]

18. Bowman,  T.E.  1977.  Isopod   crustaceans  (except Anthuridae) collected on the Presidential  Cruise  of 1938. Proc. Biol. Soc. Washington 89: 653-666.         [ Links ]

19. Bradbury, M.G. 1980. A revision of the  fish genus Ogcocephalus with descriptions of new species from the  western Atlantic Ocean (Ogcocephalidae:  Lophiformes). Proc. California Acad. Sci. 42: 229-285.         [ Links ]

20. Breedy, O. 2009. Octocorals. Text: 161-167, Species list, CD: 108-111. In:  I.S. Wehrtmann & J. Cortés (Eds.).  Marine Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

21. Breedy, O. & J. Cortés. 2008. Octocorals (Coelenterata: Anthozoa: Octocorallia) of Isla del Coco, Costa Rica. Rev. Biol. Trop. 56 (Suppl. 2): 71-77.         [ Links ]

22. Breedy, O. & J. Cortés. 2011. Morphology and taxonomy of a new  species of Leptogorgia (Cnidaria: Octocorallia: Gorgoniidae) in Cocos Island National Park, Pacific Costa Rica. Proc. Biol. Soc. Washington 124: 62-69.         [ Links ]

23. Breedy, O. & H.M. Guzman. 2003. Octocorals from Costa Rica: The genus  Pacifigorgia (Coelenterata: Octocorallia: Gorgoniidae). Zootaxa 281: 1-60.         [ Links ]

23a. Breedy, O., L.P. van Ofwegen & S. Vargas. 2012. A new family of soft  corals (Anthozoa, Octocorallia, Alcyonacea) from the aphotic tropical eastern Pacific waters revealed by integrative taxonomy. Syst. Biod. 10: 351-359.         [ Links ]

24. Briggs, J.C. 1955. A monograph of the  clingfishes (Order xenopterygii). Stanford Ichthyol.  Bull.  6: 1-224.         [ Links ]

25. Brusca, R.C. & E.W. Iverson. 1985. A guide to the marine isopod crustacea of  Pacific Costa Rica. Rev. Biol. Trop. 33 (Suppl. 1): 1-77.         [ Links ]

26. Brusca, R.C. & I.S. Wehrtmann. 2009. Isopods. Text: 257-264, Species list, CD: 206-211. In:  I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

27. Bussing, W.A. 1983. A new tropical  eastern Pacific labrid fish, Halichoeres  discolor endemic to Isla del Coco. Rev. Biol. Trop. 31: 19-23.         [ Links ]

28. Bussing, W.A. 1985. Los peces de la Familia Labridae de la costa Pacífica  de  Costa Rica. Rev. Biol. Trop. 33: 81-98.         [ Links ]

29. Bussing, W.A. 1990. New species of  gobiid fishes of the genera Lythrypnus, Elacatinus and Chriolepis. Rev. Biol. Trop. 38: 99-118.         [ Links ]

30. Bussing, W.A. 1991. A new genus and two new species of tripterygiid fishes from Costa Rica. Rev. Biol. Trop. 39: 77-85.         [ Links ]

31. Bussing, W.A. 1991. A new species of eastern Pacific moray eel (Pisces: Muraenidae). Rev. Biol Trop. 39: 97-102.         [ Links ]

32. Bussing, W.A. 1997. Chriolepis  atrimelum (Gobiidae) a new species of gobiid fish from Isla del Coco, Costa Rica. Rev. Biol. Trop. 45: 1547-1552.         [ Links ]

33. Bussing, W.A. 2010. A new fish, Peristedion nesium (Scorpaeniformes: Peristediidae) from Isla del Coco, Costa Rica. Rev. Biol. Trop. 58: 1149-1156.         [ Links ]

34. Bussing, W.A. & M.I. López. 2005. Peces de la Isla del Coco y peces arrecifales de la costa Pacífica de América Central Meridional / Fishes of Cocos Island and Reef Fishes of the Pacific Coast of Lower Central America. Rev. Biol. Trop. 53 (Suppl. 2): 192 p.         [ Links ]

35. Bussing, W.A. & M.I. López. 2009.  Text: 453-458, Species list, CD:  412-473.  In: I.S. Wehrtmann and J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

36. Cairns, S.D. 1991a. A revision of the  ahermatypic Scleractinia  of  the  Galápagos  and  Cocos  Islands. Smithsonian Contr. Zool. 504. 32 p.         [ Links ]

37. Cairns, S.D. 1991b. New records of  Stylasteridae (Hydrozoa: Hydroida) from the Galápagos and Cocos Islands. Proc. Biol. Soc. Washington 104: 209-228.         [ Links ]

38. Camacho-García, Y.E. 2009. Benthic opisthobranchs. Text: 371-386, Species list, CD:  330-347. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

39.  Castellanos,  I.,  E.  Suárez-Morales  &  A.  Morales-Ramírez. 2009a. Euphausiids. Text: 199-207, Species list,  CD: 164-166. In: I.S. Wehrtmann & J.  Cortés (Eds.). Marine Biodiversity of  Costa Rica, Central America. Springer, Berlin.         [ Links ]

40. Castellanos,  I.A.,  A.  Morales-Ramírez  &  E.  Suárez-Morales.  2009b.  Appendicularians  (Urochordata). Text:  445-452,  Species  list,  CD:  411.  In:  I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

41. Castellanos-Osorio,  I.,  R.M.  Hernández-Flores,  Á. Morales-Ramírez & M. Corrales-Ugalde.   2012. Apendicularias (Urochordata) y  quetognatos (Chaetognatha) del Parque  Nacional Isla del Coco, Costa Rica. Rev. Biol. Trop. 60 (Suppl. 3): 243-255.         [ Links ]

42. Clark, H.L. 1940. Eastern Pacific  Expeditions  of the  New York  Zoological Society. XXI Notes on Echinodermas from west coast of  Central America. Zoologica 25: 331-352.         [ Links ]

43.  Clark, H.L. 1948. A report on the Echini of the warmer eastern Pacific, based on  the collections of the Velero III. Allan Hancock Pac. Exp. 8: 225-352.         [ Links ]

Cortés,  J. 1986. Biogeografía de corales  hermatípicos: el istmo centroamericano. Anales Inst. Cienc. Mar Limnol., UNAM 13: 297-304.         [ Links ]

44.  Cortés, J. 1996/1997. Biodiversidad marina de Costa Rica: Filo Cnidaria. Rev. Biol. Trop.  44(3)/45(1): 323-334.         [ Links ]

Cortés, J. 2008. Historia de la investigación  marina de la Isla del Coco, Costa Rica. Rev. Biol. Trop. 56 (Suppl. 2): 1-18.         [ Links ]

45. Cortés, J. 2009b. Zoanthids, actiniarians, and corallimorphs. Text: 157-159, Species list, CD: 105-107. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

46.  Cortés, J. 2009c. Stony corals. Text: 169-173, Species list, CD: 112-118. In:  I.S. Wehrtmann & J. Cortés (Eds.).  Marine Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

Cortés,  J. 2009d. Other taxonomic groups  (Fungi, kinorhynchs, invertebrate chordates). Text: 497-500, Species list, CD: 491-492. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of  Costa Rica, Central America. Springer, Berlin.         [ Links ]

Cortés, J. 2009e. Marine fish parasites. Text: 501-505, Species list, CD: 493-496. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine  Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

Cortés,  J. 2011. Eastern Tropical Pacific coral  reefs, Pp. 351-358.  In: D. Hopley (Ed.). The  Encyclopedia of Modern Coral Reefs:  Structure, Form and Process. Springer, Berlín.         [ Links ]

Cortés, J. 2013a. Cocos Island marine  ecosystems.  In M.  Kappelle  (Ed.).  Costa  Rican  Ecosystems.  The University  of Chicago Press, Chicago and London. In press.         [ Links ]

Cortés, J. 2013b. The Pacific coastal and marine ecosystems. In M. Kappelle (Ed.). Costa Rican  Ecosystems. The University of Chicago  Press, Chicago and London. In press.         [ Links ]

47.  Cortés, J. & S. Blum. 2008. Life down to 450 m at Isla del Coco, Costa Rica. Rev. Biol. Trop. 56 (Suppl. 2): 189-206.         [ Links ]

48.  Cortés, J. & H.M. Guzmán. 1998. Organismos de los arrecifes coralinos de  Costa Rica: Descripción, distribución geográfica e historia natural de los corales zooxantelados (Anthozoa: Scleractinia) del Pacífico. Rev. Biol. Trop. 46: 55-92.         [ Links ]

Cortés,  J. & E. Salas. 2009. Seagrasses.  Text: 119-122, Species list, CD: 71-72. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

Cortés,  J. & I.S. Wehrtmann. 2009. Diversity  of marine habitats of the Caribbean and Pacific of Costa Rica. Capítulo I, Pp. 1-45. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa  Rica, Central America. Springer, Berlin.         [ Links ]

49. Cortés,  J.,  C.  Mora-Baumgartner  & V.   Nielsen. 2009a.  Foraminiferans. Text: 131-135,  Species list, CD: 79-82. In: I.S.  Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

50.  Cortés,  J.,  N. van der  Hal  &  R.W.M.  van Soest. 2009b.  Sponges.  Text:  137-142,  Species  list,  CD: 83-93. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

51. Cortés, J., v.  Nielsen & A.  Herrera-Cubilla. 2009c. Bryozoans. Text: 413-416, Species list, CD: 385-388. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa  Rica, Central America. Springer, Berlin.         [ Links ]

52. Cortés, J., A. Sánchez-Jiménez, A. Rodríguez-Arrieta, G. Quirós-Barrantes, P.C. González & S. Blum. 2012. Elasmobranchs observed in deep waters (45-330m) at Isla del Coco National Park, Costa Rica (Eastern Tropical Pacific). Rev. Biol. Trop. 60 (Suppl. 3): 257-273.         [ Links ]

53. Crane, J. 1975. Fiddler Crabs of the World. Ocypodidae: Genus Uca.  Princeton Univ. Press, Princeton, New Jersey. 738 p.         [ Links ]

54.  Crouch, R.W. & C.W. Poag. 1987. Benthic Foraminifera of the Panamanian Province: distribution and origins. J. Foram. Res. 17: 153-176.         [ Links ]

55.  Cushman, J.A. & I. McCulloch. 1939. A report on some arenaceous  Foraminifera. Allan Hancock Pac. Exped. 6: 1-113.         [ Links ]

56.  Cushman,  J.A.  &  I.  McCulloch.  1942.  Some Virgulininae in the Collections  of  the Allan Hancock Foundation. Allan Hancock Pac. Exped. 6: 179-230.         [ Links ]

57.  Cushman, J.A. & I. McCulloch. 1950. Some Lagenidae in the collections of the Allan Hancock Foundation. Allan Hancock Pac. Exped. 6: 295-364.         [ Links ]

58. Cutler,  N.,  E.  Cutler  &  J.A. Vargas.  1992.  Peanut worms  (Phylum  Sipuncula)  from  Costa  Rica.  Rev. Biol. Trop. 40: 153-158.         [ Links ]

59.  Dall, W.H. 1895. Scientific results of explorations by the  U.S.  Fish  Commission  Steamer Albatross.  No. XXXIV. Report on Mollusca and  Brachiopoda dredged in deep water, chiefly near the Hawaiian Islands, with  illustrations of hitherto unfigured species  from northwest America. Proc. U.S. Natl.  Mus. 17(1032): 675-733.         [ Links ]

60. Dall, W.H. 1908. Reports on the dredging operations off the west coasts of  Central America to the Galapagos, to  the  west coast of Mexico, and in the  Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission steamer “Albatross”, during 1891, Lieut. Commander Z.L. Tanner, U.S.N., commanding. XXXVII, and Reports on the scientific results of the  expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer “Albatross,” from October, 1904, to  March, 1905, Lieut. Commander L.M.  Garrett, U.S.N., commanding. XIV. The Mollusca and Brachiopoda. Bull. Mus.  Comp. Zoöl., Harvard Coll. XLIII: 205-487.         [ Links ]

61. D’Attilio, A. & B.W. Myers. 1988. A new species of Favartia from the eastern Pacific (Gastropoda: Muricidae). Nautilus 102: 106-109.         [ Links ]

62. Dawson, C.E. 1977. Studies of the  eastern Pacific sand stargazers (Pisces:  Dactyloscopidae). 4. Gillellus,  Sindoscopus new genus, and Heteristius  with description of new species. Proc.  California Acad. Sci. 41: 125-160.         [ Links ]

De la Cruz, E.M. & J.A. Vargas. 1986. Estudio preliminar de la meiofauna de la playa fangosa de Punta Morales, Golfo de  Nicoya, Costa Rica. Brenesia 25-26: 89-87.         [ Links ]

63.  Dean, H.K. 2004. Marine biodiversity of Costa Rica: Class  Polychaeta  (Annelida).  Rev.  Biol.  Trop.  52 (Suppl. 2): 131-181.         [ Links ]

64.  Dean, H.K. 2009. Polychaetes and  echiurans. Text: 181-191,  Species list, CD: 122-159. In:  I.S. Wehrtmann  &  J.  Cortés  (Eds.).  Marine  Biodiversity  of Costa Rica, Central America. Springer, Berlin.         [ Links ]

65. Dean, H.K., J.A. Sibaja-Cordero, J. Cortés, R. Vargas & G.Y. Kawauchi. 2010a. Sipunculids and Echiurans of Isla del Coco (Cocos Island), Costa Rica. Zootaxa 2557: 60-68.         [ Links ]

66. Dean, H.K., J.A. Sibaja-Cordero & J. Cortés. 2010b. Occurrence of the phoronid Phoronopsis albomaculata in Cocos Island, Costa Rica. Pac. Sci. 64: 459-462.         [ Links ]

67.  Dean, H.K., J.A. Sibaja-Cordero & J.  Cortés. 2012. Polychaetes (Annelida: Polychaeta) of Parque Nacional Isla del Coco, Pacific  Costa Rica. Pac. Sci. 63: 347-386.         [ Links ]

Dexter, D.M. 1974. Sandy-beach fauna of the  Pacific and Atlantic coasts of Costa Rica and Colombia. Rev. Biol. Trop. 22: 51-66.         [ Links ]

68. Díaz-Agras, G. 2008. Revision of the genus Pozziella (Porifera:  Poecilosclerida)  with  description  of  three new species  from the eastern Pacific. Zootaxa 1866: 69-94.         [ Links ]

69. Emerson, W.K. 1994. A zoogeographic summary of the marine mollusks of Clipperton Island (Tropical Eastern Pacific Ocean). Festivus 26: 62-71.         [ Links ]

70. Emerson, W.K. 1995. A zoogeographic summary of the marine mollusks of the Revillagigedo Islands (Tropical Eastern Pacific Ocean). Festivus 27: 3-18.         [ Links ]

71. Emig, C.C. 2009. Brachipods. Text: 417-420, Species list, CD: 389-391. In:  I.S. Wehrtmann & J. Cortés (Eds.).  Marine Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

72. Excoffon, A.C., F.H. Acuña & J. Cortés.  2009. The sea anemone Nemanthus californicus (Cnidaria, Actiniaria, Nemanthidae) from Costa Rica: re-description and first record outside the type locality. Mar. Biodiv. Rec. 2: 1-5.         [ Links ]

73. Faxon, W. 1893. Reports on the dredging operations off the west coasts of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission  steamer “Albatross”, during 1891, Lieut. Commander Z.L. Tanner, U.S.N., commanding. VI. Preliminary  descriptions  of  new species of Crustacea. Bull. Mus. Comp. Zoöl., Harvard Coll. XXIV: 149-220.         [ Links ]

74. Fernández, C. 2008. Flora marina del Parque Nacional  Isla  del  Coco,  Costa  Rica,  Pacífico  Tropical Oriental. Rev. Biol. Trop. 56 (Suppl. 2): 57-69.         [ Links ]

75. Fernández, C. & J.J. Alvarado. 2008. Chlorophyta de la costa Pacífica de Costa Rica. Rev. Biol. Trop. 56 (Suppl. 4): 149-162.         [ Links ]

76. Fernández-García,  C.,  R.  Riosmena-Rodríguez,  B. Wysor, O.L. Tejada & J. Cortés. 2011. Checklist of the Pacific marine  macroalgae of Central America. Bot. Mar. 54: 53-73.         [ Links ]

77. Fernández-Leiva, S. 1996. Taxonomía del ictioplancton en la Isla del Coco,  Costa  Rica. Thesis, Univ. Costa Rica, San Pedro, Costa Rica. 175 p.         [ Links ]

78. Ferreira, A.J. 1987. The chiton fauna of Cocos Island, Costa Rica (Mollusca: Polyplacophora) with the description of two new species. Bull. Sci. California Acad. Sci. 86: 41-53.         [ Links ]

79. Foster, J.M., S.E. Lecroy, R.W. Heard &  R. Vargas. 2009. Gammaridean Amphipods. Text 265-274, Species list, CD: 212-216. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine  Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

80. Fraser, C.M. 1938. Hydroids of the 1932, 1933, 1935, and 1938 Allan Hancock  Pacific Expeditions. Allan Hancock Pac. Exped. 4: 129-153.         [ Links ]

81. Fraser, C.M. 1948. Hydroids of the  Allan Hancock Pacific Expeditions since March, 1938. Allan Hancock Pac. Exped. 4: 179-335.         [ Links ]

82. Gallardo, V.A. & C. Espinoza. 2007. New communities of large filamentous sulfur bacteria in the eastern South Pacific. Int. Microbiol. 10: 97-102.         [ Links ]

83. Garman, S. 1899. Reports on an  exploration off the west coasts of  Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer “Albatross”, during  1891, Lieut. Commander Z.L. Tanner U.S.N., commanding. XXVI. The Fishes. Mem. Mus. Comp. Zoöl., Harvard Coll. XXIV: 1-432.         [ Links ]

84. Garrison, G. 2005. Peces de la Isla del Coco/Isla del Coco Fishes, 2nd ed. Edit.  INBio, Heredia, Costa Rica. 429 p.         [ Links ]

85. Garth, J.S. 1991. Taxonomy,  distribution, and ecology of Galápagos  Brachyra. Pp. 123-145. In: M.J. James (Ed.). Galápagos Marine Invertebrates: Taxonomy,  Biogeography,  and   Evolution  in  Darwin’s Islands. Plenum Press, New York, London.         [ Links ]

86. Gasca, R. 2009. Hyperiid amphipods. Text: 275-282. Species  list, CD: 217-218. In: I.S. Wehrtmann & J. Cortés  (Eds.).  Marine  Biodiversity  of  Costa  Rica, Central America. Springer, Berlin.         [ Links ]

87. Gasca,  R.  &  Á.  Morales-Ramírez.  2012.  Anfípodos  hiperídeos  (Crustacea: Peracarida) del Parque Nacional Isla del Coco, Costa Rica, Pacífico Tropical Oriental. Rev. Biol. Trop. 60 (Suppl. 3): 223-233.         [ Links ]

Glynn, P.W. & J. Ault. 2000. A biogeographic analysis and review of the far eastern  Pacific coral reef region. Coral Reefs 19: 1-23.         [ Links ]

88. Gore, R.H. 1982. Porcellanid crabs from the coasts of Mexico and Central America (Crustacea: Decapoda: Anomura). Smithsonian Contr. Zool. 363: 1-34.         [ Links ]

89. Grau, G. 1959. Pectinidae of the eastern Pacific. Allan Hancock Pac. Exped. 23: 1-308.         [ Links ]

90. Haig, J. 1960. The Porcellanidae (Crustacea: Anomura) of the eastern Pacific. Allan Hancock Pac. Exped. 24: 1-440.         [ Links ]

91. Haig, J. & A.J. Provenzano. 1965. A new genus and two new species of diogenid hermit crabs (Decapoda, Anomura). Crustaceana 9: 199-207.         [ Links ]

92. Hartman,  O.  1939a.  Polychaetous  Annelids.  Part  I. Aphroditidae to Pisionidae. Allan Hancock Pac. Exped. 7: 1-155.         [ Links ]

93. Hartman, O. 1939b. The Polychaetous Annelids collected on the Presidential  Cruise of 1938. Smithsonian Miscel. Coll. 98(13): 1-22.         [ Links ]

94. Hartman,  O.  1940.  Polychaetous  Annelids.  Part  II. Chrysopetalidae to Goniadidae. Allan  Hancock Pac. Exped. 7: 173-287.         [ Links ]

95. Hastings, P.A. & D.R. Robertson 1999. Acanthemblemaria atrata and  Acanthemblemaria mangognatha, new  species  of  eastern  Pacific  barnacle  blennies (Chaenopsidae) from Isla del Coco, Costa Rica, and Islas  Revillagigedo, Mexico, and their  relationships with other barnacle  blennies. Rev. Francaise Aquariol. Herpetol. 25: 107-118.         [ Links ]

96. Heard, R.W., O. Breedy & R. Vargas. 2009. Tanaidaceans. Text: 245-256, Species list, CD: 204-205. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

97. Heller, E. & R.E. Snodgrass. 1903. Papers from the Hopkins-Stanford Galapagos Expedition 1898-1899. XV.  New  fishes.  Proc.  Washington  Acad.  Sci.  5: 189-229.         [ Links ]

98. Hertlein, L.G. 1963. Contribution to the biogeography of Cocos Island, including a bibliography. Proc. California Acad. Sci. 4th Ser. 32: 219-289.         [ Links ]

Heiner, I. & B. Neuhaus. 2007. Loricifera from the deep sea at the Galápagos Spreading Center, with a description of Spinoloricus turbatio gen. et sp. nov. (Nanaloricidae). Helgol. Mar. Res. 61: 167-182.         [ Links ]

Hickman, C.P., Jr. 2009. Evolutionary responses of marine invertebrates to insular isolation in Galapagos. Galápagos Res. 66: 32-42.         [ Links ]

99. Hochberg, F.G. & Y.E. Camacho-García. 2009. Squids and octopuses. Text: 399-407, Species list, CD: 379-382. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

100. Hollmann, M. 1996. Polinices (Mammilla) simiae on Isla del Coco, Costa Rica: another Indo-Pacific invader into the Panamic Province. Festivus 28: 24-29.         [ Links ]

101.  Howard, A.D. 1952. Pteropods collected by the Allan Hancock Foundation. Minutes of the Conchological Club of Southern California, No. 121, pp. 12-14.         [ Links ]

102.  Hoyle, W.E. 1904. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U. S. Fish  Commission steamer “Albatross,” during 1891, Lieut. Commander Z. L. Tanner, U.S.N., Commanding, XXIX. Reports on the Scientific results of the expedition to the tropical Pacific, in charge of Alexander Agassiz, on the U. S. Fish  Commission steamer “Albatross,” from  August, 1899, to March,V1900, Commander Jefferson T. Moser, U. S. N., commanding, V. Cephalopoda. Bull.  Mus. Comp. Zoöl., Harvard Coll. XLIII: 1-71.         [ Links ]

103. Hutchings, P., A. Reid, & R. Wilson.  1991. Perinereis (Polychaeta,  Nereidiae)  from Australia, with redescriptions of six additional species. Rec. Australian Mus. 43: 241-274.         [ Links ]

104. Iwamoto, T. & Y.I. Sazonov. 1988. A review of the southeastern  Pacific   Coryphaenoides  (sensu  lato) (Pisces, Gadiformes, Macrouridae). Proc. California Acad. Sci. 45: 35-82.         [ Links ]

105. Iwamoto,  T.  &  J.E.  McCosker.  2001.   Notes  on Galápagos grenadiers (Pisces, Gadiformes, Macrouridae),  with  the  description of a new species of Coryphaenoides. Rev. Biol. Trop. 49  (Suppl.  1): 21-27.         [ Links ]

106. Jiménez-Cueto, S., E. Suárez-Morales & Á. Morales-Ramírez. 2012. Algunos poliquetos holoplanctónicos (Annelida:  Polychaeta) del Parque Nacional de Isla del Coco, Costa Rica. Rev. Biol. Trop. 60 (Suppl. 3): 207-222.         [ Links ]

107. Kabat, A.R. 2000. Results of the Rumphius Biohistorical Expedition to Ambon (1990). Part 10. Mollusca,  Gastropoda,  Naticidae.  Zool.  Med.  Leiden  73: 345-380.         [ Links ]

108. Kaiser, K.L. 2001. Comments on four  Muricoidean (Mollusca) species formerly endemic to Isla del Coco found at Isla de Malpelo. Festivus 33: 3-7.         [ Links ]

109. Kelmo, F. & R. Vargas. 2002.  Anthoathecatae and Leptothecatae  hydroids from Costa Rica (Cnidaria: Hydrozoa). Rev. Biol. Trop. 50: 599-627.         [ Links ]

Knowlton, N. 2000. Molecular genetic analyses of species boundaries in the sea. Hydrobiologia 420: 73-90.         [ Links ]

110. Laguarda-Figueras  A.  &  F.A.   Solís-Marín.  2009. Holothuria (Cystipus) casoae a new  species  of sea cucumber   (Echinodermata:   Holothuroidea)   from the central-eastern Pacific Ocean. Scient.  Mar.  73: 573-578.         [ Links ]

111. Lalicker, C.G. & I. McCulloch. 1940. Some Textulariidae of the Pacific Ocean. Allan Hancock Pac. Exped. 6: 115-143.         [ Links ]

112. Lattig, P. & S.D. Cairns. 2000. A new species of Tethocyathus (Cnidaria: Anthozoa: Scleractinia: Caryophylliidae), a trans-isthmian azooxanthellate species. Proc. Biol. Soc. Washington 113: 590-595.         [ Links ]

Lessios, H.A. & D.R. Robertson. 2006. Crossing the impassable:  genetic  connections  in  20  reef  fishes  across the  Eastern  Pacific  Barrier.  Proc.  R.  Soc.  Lond  B 273:.2201-2208.         [ Links ]

113. Lessios, H.A., B.D. Kessing & D.R. Robertson. 1998. Massive  gene  flow  across  the  world’s  most  potent marine biogeographic barrier.  Proc. R. Soc. Lond. B 265:583-588.         [ Links ]

114. Lindberg, D.R. & J.H. McLean. 1981. Tropical Eastern Pacific limpets of the  Family Acmaeidae (Mollusca, Archaeogastropoda): Generic criteria and  description of six new species from the mainland and the Galápagos Islands. Proc. California Acad. Sci. 42: 323-339.         [ Links ]

Lizano, O.G. 2008. Dinámica de aguas alrededor de la Isla del Coco, Costa Rica.  Rev. Biol. Trop. 56 (Supl. 2): 31-48.         [ Links ]


115. Long, D.J., J.E. McCosker, S. Blum &  A. Klapfer. 2011. Tropical Eastern Pacific Records of the Prickly Shark, Echinorhinus cookei (Chondrichthyes: Echinorhinidae). Pac. Sci. 65: 433-440.         [ Links ]

116. López-Garro, A., I. Zanella, G. Golfín-Duarte & M. Pérez-Montero. 2012. First record of the blacktip reef shark Carcharhinus melanopterus (Quoy & Gaimard, 1824) (Carcharhiniformes: Carcharhinidae) from the Eastern Tropical Pacific. Rev. Biol. Trop. 60 (Suppl. 3): 275-278.         [ Links ]

117. Ludwig, H.L. 1905. Reports on an exploration off the West Coast of Mexico,  Central and South America, and off the Galapagos islands, in charge of Alexander Agassiz, by the U. S. Fish Commision steamer “Albatros”, during 1891. XXXV. Reports on the Scientific results of the  expedition to the Tropical Pacific in Charge of A. Agassiz on the Fish Commision Steamer “Albatros”, from August, 1899, to March, 1900. VII. Asteroidea. Mem. Mus. Comp. Zoöl., Harvard Coll. XXXII: 1-292.         [ Links ]

118. Magaña-Cubillo, J.A. & J. Espinosa. 2009. Bivalves. Text: 387-398, Species list, CD:  348-378. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

119. Maluf, L.Y. 1991. Echinoderm fauna of the Galápagos  Islands.  Pp.  345-367.  In:  M.  J.  James  (Ed.). Galápagos Marine Invertebrates: Taxonomy, Biogeography, and Evolution in Darwin’s  Islands. Plenum Press, New York and London.         [ Links ]

Manning,  R.B. 1972. Three new stomatopod  crustaceans of the Family Lysiosquillidae from the eastern Pacific Region. Proc. Biol. Soc. Washington 85: 271-278.         [ Links ]

120.  May-Collado, L. 2009. Marine mammals. Text: 479-495, Species list, CD: 485-490. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

121. McCosker,  J.E.  1988. A revision  of  the  snake-eel genus  Callechelys   (Anguilliformes:  Ophichthidae) with the description of two new Indo-Pacific species and a new callechelyin genus. Proc. California Acad. Sci. 50: 185-215.         [ Links ]

122. McCosker, J.E. & R.H. Rosenblatt. 2010. The fishes of the Galápagos archipelago: an update. Proc. California Acad. Sci, Ser. 4, 61 (Suppl. II): 167-195.         [ Links ]

123. McCosker, J.E.,  J.S.  Stephens  &  R.H.  Rosenblatt. 2003.  Cottoclinus canops, a new genus  and species of  blenny  (Perciformes:   Labrisomidae)  from  the Galápagos  Islands. Proc. California Acad. Sci. 54: 155-160.         [ Links ]

124. McLaughlin,  P.A.  1982.  Revision  of  Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species: Part III. Agaricochirus McLaughlin, and Enallopaguropsis McLaughlin. Bull. Mar. Sci. 32: 823-856.         [ Links ]

125. McLean, J.H. 1969. New species of tropical eastern Pacific Gastropoda. Malac. Rev. 2: 115-130.         [ Links ]

126. Meisler, M.R. & R.J. Lavenberg. 1998. A new species (Serranidae: Serranus) from Isla del Coco, Pp. 115. In: G.R. Allen &  D.R. Robertson (Eds.). Peces del Pacífico  Oriental Tropical.  Crawford  House  Press, Bathhurst, Australia.         [ Links ]

127. Møller, P.R., W. Schwarzhans & J.G. Nielsen. 2005. Review  of the American Dinematichyini  (Teleostei: Bythitidae). Part II. Ogilbia. Aqua 10: 133-207.         [ Links ]

128. Montoya,  M.  2008a. Aves  marinas  de  la  Isla  del Coco, Costa Rica, y su conservación. Rev. Biol. Trop. 56 (Suppl. 2): 133-149.         [ Links ]

129. Montoya, M. 2008b. La presencia de otáridos (Carnivora: Otariidae) en la isla del Coco, Costa Rica. Rev. Biol. Trop. 56 (Suppl. 2): 151-158.         [ Links ]

Montoya, M. & K.L. Kaiser. 1988. Biogeographical notes on the genus Terebra (Gastropoda) at Isla del Coco, Costa Rica. Rev. Biol. Trop. 36: 569-574.         [ Links ]

130. Morales-Ramírez, A. 2008. Caracterización cualitativa del zooplancton del Área de Conservación Marina Isla del Coco (ACMIC), Océano  Pacífico de Costa Rica. Rev. Biol. Trop. 56 (Suppl. 2): 159-169.         [ Links ]

131. Morales-Ramírez, A.  &  E.  Suárez-Morales.  2009. Copepods. Text: 291-305, Species list, CD: 224-236. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa  Rica, Central America. Springer, Berlin.         [ Links ]

132. Motomura, H. 2004. New species of  scorpionfish, Scorpaena  cocosensis   (Scorpaeniformes:  Scorpaenidae) from  the Cocos Islands, Costa Rica, eastern Pacific Ocean. Copeia 2004: 818-824.         [ Links ]

Neuhaus,  B. 2004. Description of  Campyloderes cf. vanhoeffeni (Kinorhyncha, Cyclorhagida) from the Central American East Pacific Deep with a review of the genus. Meiofauna Mar. 13: 3-20.         [ Links ]

Neuhaus,  B.  &  T.  Blasche.  2006.  Fissuroderes,  a  new genus of Kinorhyncha (Cyclorhagida) from the deep sea and continental shelf of New Zealand and from the continental shelf of Costa Rica. Zool. Anz. 245: 19-52.         [ Links ]

Nova-Bustos,  N., A.C.  Hernández-Zanuy  &  R. Viquez-Portuguez. 2010. Distribución y  abundancia de las ascidias de los fondos rocosos de la Bahía de Cuajiniquil, Costa Rica. Bol. Invest. Mar. Cost. 39: 57-66.         [ Links ]

133.  Osburn, R.C. 1950. Bryozoa of the Pacific Coast of America. Part 1, Cheilostomata-Anasca. Allan Hancock Pac. Exped. 14: 1-269.         [ Links ]

134. Osburn, R.C. 1952. Bryozoa of the Pacific Coast of America.  Part  2,  Cheilostomata-Ascophora.  Allan Hancock Pac. Exped. 14: 271-611.         [ Links ]

135. Osburn, R.C. 1953. Bryozoa of the Pacific Coast of America. Part 3, Cyclostomata, Ctenostomata, Entoprocta, and Addenda. Allan Hancock Pac. Exped. 14: 613-725.         [ Links ]

136. Perger, R., R. vargas & A. Wall. 2011. Johngarthia cocoensis, a new species of Gecarcinidae MacLeay, 1838 (Crustacea, Decapoda, Brachyura) from Cocos Island, Costa Rica. Zootaxa 2911: 57-68.         [ Links ]

Petrescu,  I., R.W. Heard, R. Vargas  & O.  Breedy. 2009. Cumaceans.  Text: 237-244, Species list,  CD: 201-203. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

137. Petuch,  E.J.  &  D.M.  Sargent.  1986.  Atlas  of  the Living Olive Shells of the World. Coastal Education and Research Foundation, Jacksonville. 253 p.         [ Links ]

Polidoro, B.A., T. Brooks, K.E. Carpenter, G.J. Edgar, S. Henderson,  J. Sanciangco & D.R.  Robertson. 2012. Patterns of extinction risk and threat for marine vertebrates  and  habitat-forming species in the Tropical Eastern Pacific. Mar. Ecol. Prog. Ser. 448: 93-104.         [ Links ]

138. Poss, S.G., J.E. McCosker & C.C. Baldwin. 2010. A new species of Scorpaenodes (Pisces: Scorpaenidae) from  the Galápagos and Cocos islands with  discussions  of  the  limits  of  Scorpaenodes  and  Thysanichthys. Proc. California Acad. Sci. 61: 235-267.         [ Links ]

139. Price, W.W., R.W. Heard & R. Vargas. 2009. Shallow water mysids. Text: 229-236, Species list, CD: 199-200. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

140. Randall, J.E. & J.E. McCosker. 1975. The eels of Easter Island with a description of a new moray. Contrib. Sci., Nat. Hist. Mus. Los Angeles County 264: 1-32.         [ Links ]

141. Reiswig, H.M. 2010. A new species of Tretodictyum (Porifera:  Hexactinellida:  Tretodictyidae)  from  off Cocos Island,  tropical eastern Pacific Ocean. Proc. Biol. Soc. Washington 123: 242-250.         [ Links ]

142. Robertson,  D.R.  &  G.R. Allen.  2008.  Shorefishes of  the Tropical  Eastern  Pacific online  information system. Version 1.0. Smithsonian Tropical Research Institute,  Balboa,  Panamá.   www.neotropicalfishes. org/sftep, www.stri.org/sftep         [ Links ]

Robertson,  D.R. & K.L. Cramer. 2009. Shore  fishes and biogeographic subdivisions of  the Tropical Eastern Pacific. Mar. Ecol. Prog. Ser. 380: 1-17.         [ Links ]

143. Robertson, D.R., J.S. Grove & J.E. McCosker. 2004. Tropical  transpacific  shore  fishes.  Pac.   Sci.  58: 507-565.         [ Links ]

144. Robson,  G.C.  1929.  A  monograph  of  the  Recent Cephalopoda based on the collections in  the  British Museum (Natural History). Part 1. Octopodinae. Printed by order of the Trustees of the British Museum, July 27: 1-236.         [ Links ]

145. Robson,  G.C.  1932.  A  monograph  of  the  Recent Cephalopoda based on the  collections in the British Museum  (Natural  History). Part 2. Octopoda.  Printed by order of the Trustees of the  British Museum, January 23: 1-359.         [ Links ]

146.  Robson, G.C. 1948. The Cephalopoda Decapoda of the Arcturus Oceanographic Expedition, 1925. Zoologica 33: 115-132.         [ Links ]

147.  Rodríguez-Saénz, K. 2005. Distribución  espacial y temporal de la biomasa,  composición y abundancia del  zooplancton,  con  énfasis  en  hidromedusas  de Bahía Culebra durante  La Niña 1999-2000. M.Sc. Tesis,  Univ.  Costa  Rica,  San  Pedro,  Costa  Rica. 156 p.         [ Links ]

148. Rodríguez-Sáenz, K. & R. Gasca. 2009. Siphonophores. Text: 151-156, Species list, CD: 101-104. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

149. Rodríguez-Sáenz,  K.  &  L.  Segura-Puertas.  2009. Hydrozoa, Scyphozoa and Cubozoa  (Medusozoa). Text: 143-149, Species  list,  CD: 143-149. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

150. Rodríguez-Sevilla, L., R. Vargas & J. Cortés. 2009. Benthic,  shelled gastropods. Text:  333-355,  Species list, CD: 243-325. In:  I.S. Wehrtmann & J. Cortés (Eds.).  Marine Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

151. Rosenblatt,  R.H.  &  J.E.  McCosker.  1988. A new species of  Acanthemblemaria from Malpelo Island, with a key to the Pacific members of the genus (Pisces: Chaenopsidae). Proc.  California Acad. Sci. 45: 103-110.         [ Links ]

152. Roth, B. & E. V. Coan. 1971. Marginellidae (Mollusca: Neogastropoda) from the Galápagos Islands and Cocos Island. Proc. California Acad. Sci. 4th Ser. 37: 575-584.         [ Links ]

153. Rusch, D.B. + 39 more authors. 2007. The Sorcerer II Global Ocean Sampling Expedition: northwest Atlantic through eastern tropical Pacific. PloS Biol. 5(3): e77. doi:101371/journal.pbio.0050077         [ Links ]

Santoro,  M. & S. Mattiucci. 2009. Sea turtle  parasites. Text: 507-519, Species list, CD:  497-500.  In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

154. Sasa, M., G.A. Chaves & L.D. Patrick. 2009. Marine reptiles and amphibians. Text: 459-468, Species list, CD: 474-478. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

155. Schultz, L.P. 1944. A revision of the  American clingfishes, Familia Gobiesocidae, with descriptions. of new genera and forms. Proc. U.S. Nat. Mus. 96: 47-77.         [ Links ]

156. Schwabe, E. & I.S. Wehrtmann. 2009. Chitons. Text: 323-331,  Species list, CD: 240-242. In:  I.S. Wehrtmann  &  J.  Cortés  (Eds.).  Marine  Biodiversity  of Costa Rica, Central America. Springer, Berlin.         [ Links ]

157. Shoemaker,  C.R.  1942. Amphipod   crustaceans collected on the Presidential Cruise of 1938. Smithsonian Misc. Coll. 101: 1-52.         [ Links ]

158. Sibaja-Cordero, J.A., J. Troncoso & J. Cortés. 2012. The lancelet Asymmetron lucayanum   complex  in Cocos Island National Park, Pacific Costa Rica. Pac. Sci. 66: 521-526.         [ Links ]

Silva-Benavides,  A.M. 2009. Mangroves. Text:  123-130, Species list, CD: 73-78. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

159. Solis-Marín, F.A., J.J. Alvarado +70  more authors. 2013. Appendix, p. 543-654. In J.J. Alvarado & F.A. Solís-Marín (ed.). Echinoderm Research and Diversity in Latin America. Springer, Berlin, Heidelberg.         [ Links ]

160. Solano-Barquero, A. 2011. Macrofauna  asociada a rodolitos en el Parque Nacional Isla del Coco, Costa Rica. Thesis, Escuela de Biología, Univ. Costa Rica, San Pedro, Costa Rica. 50 p.         [ Links ]

161. Solórzano, A. 2004. Serpientes de Costa  Rica: distribución, taxonomía e historia natural. Edit. INBio, Santo Domingo de Heredia, Costa Rica. 791 p.         [ Links ]

162. Springer, M. 2009.  Marine  insects.  Text:  313-322, Species list, CD: 239. In: I.S. Wehrtmann & J. Cortés (Eds.).  Marine  Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

163. Starr, R.M., K. Green & E. Sala. 2012. Deep-water fish assemblages at Coco  Island National Park and Las Gemelas Seamounts, Costa Rica. Rev. Biol. Trop. 60 (Suppl. 3): 347-362.         [ Links ]

164. Stebbing, T.R.R. 1903. Amphipoda from Costa Rica. Proc. U.S. Nat. Mus. 26: 925-931.         [ Links ]

165.  Suárez-Morales,  E.  &  R.  Gasca.   2012.  A  new Lepeophtheirus    (Copepoda: Siphonostomatoida: Caligidae) from Isla del Coco, Costa  Rica, Eastern Tropical  Pacific.  Rev.  Biol.  Trop.  60  (Suppl.  3): 235-242.         [ Links ]

166. Suárez-Morales,  E.  &  A.  Morales-Ramírez.  2001. Nuevo registro de Acartia (Planktacartia)  negligens (Copepoda, Calanoida) en el Pacífico Tropical Oriental. Rev. Biol. Trop. 49: 1286.         [ Links ]

167. Suárez-Morales,  E.  &  A.  Morales-Ramírez.  2009. New species of Monstrilloida (Crustacea: Copepoda) from the Eastern Tropical Pacific. J. Nat.  Hist. 43: 1257–1271.         [ Links ]

168. Suárez-Morales,  E.,  R.M.  Hernández-Flores  &  A. Morales-Ramírez. 2009. Chaetognaths or   arrow worms. Text: 435-443, List of species: CD  409-410. In: I.S. Wehrtmann and J. Cortés (Eds.). Marine Biodiversity of Costa  Rica, Central America. Springer, Berlín.         [ Links ]

169. Thoma, B.P., R.W. Heard, R. Vargas.  2005. A new species of Parapinnixa  (Decapoda: Brachyura: Pinnotheridae) from Isla del Coco, Costa Rica. Proc. Biol. Soc. Washington 118: 543-550.         [ Links ]

Tokioka, T. 1971. A new species of Rhopalaea from the Pacific coast of Costa Rica  (Tunicata, Ascidiacea). Publ. Seto Mar. Biol. Lab. XIX (2/3): 119-122.         [ Links ]

Tokioka, T. 1972. On a small collection of ascidians from the Pacific coast of Costa Rica. Publ. Seto Mar. Biol. Lab. XIX(6): 383-408.         [ Links ]

170. Tovar-Hernández, M.A. & H. Dean. 2010. Four new species of fan worms  (Polychaeta: Sabellidae) from worldwide localities. Scient. Mar. 74: 815-826.         [ Links ]

171. Treadwell, A.L. 1928. Polychaetous  annelids  from the Arcturus Oceanographic Expedition. Zoologica 8: 449-485.         [ Links ]

Ulken, A., R. Víquez,  C. Valiente  & M.  Campos. 1990. Marine fungi (Chytridiomycetes and  Thraustochytriales)  from  a  mangrove  area  at  Punta  Morales, Golfo de  Nicoya,  Costa Rica. Rev. Biol. Trop.  38: 243-250.         [ Links ]

172.  Valdés, Á. & Y.E. Camacho-García. 2004. “Cephalaspidean” Heterobranchs (Gastropoda) from the Pacific Coast of Costa Rica. Proc. California Acad. Sci. 55: 459-497.         [ Links ]

Van  Name, W.G. 1945. The North and South  American Ascidians. Bull. American Mus. Nat. Hist. 84: 1-476.         [ Links ]

173. Van Syoc, R. 2009. Barnacles. Text: 283-289, Species list, CD: 219-223. In:  I.S. Wehrtmann & J. Cortés (Eds.).  Marine Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

Vargas, J.A. 1988a. Community structure of macrobenthos and the results of macropredator exclusion on a tropical mud flat. Rev. Biol. Trop. 36: 287-308.         [ Links ]


Vargas, J.A. 1988b. A survey of the meiofauna of an eastern tropical Pacific intertidal  mud flat. Rev. Biol. Trop. 36: 541-544.         [ Links ]

174. Vargas, J.A. & H.K. Dean. 2009. Sipunculans. Text: 175-180,  Species list, CD: 119-121. In:  I.S. Wehrtmann  &  J.  Cortés  (Eds.).  Marine  Biodiversity  of Costa Rica, Central America. Springer, Berlin.         [ Links ]

175. Vargas, J.A. & H.K. Dean. 2010. On Branchiostoma californiense  (Cephalochordata)  from  the  Gulf  of Nicoya estuary, Costa Rica. Rev. Biol.  Trop.  58: 1143-1148.         [ Links ]

176. Vargas, R. 2009. Stomatopods. Text: 193-197, Species list, CD: 160-163. In:  I.S. Wehrtmann & J. Cortés (Eds.).  Marine Biodiversity of Costa Rica,  Central America. Springer, Berlin.         [ Links ]

177. Vargas-Castillo, R. & I.S. Werhtmann.  2008.  Stomatopods and decapods from Isla del Coco, Pacific Costa Rica. Rev. Biol. Trop. 56 (Suppl. 2): 79-97.         [ Links ]

178. Vargas, R. & I.S. Werhtmann. 2009. Decapod crustaceans. Text: 209-228, Species list, CD: 167-198. In: I.S. Wehrtmann & J. Cortés (Eds.). Marine Biodiversity of Costa Rica, Central America. Springer, Berlin.         [ Links ]

179. Vargas, R., S. Jesse & M. Castro. 1996. Checklist of crustaceans (Decapoda and  Stomatopoda), collected during  the  Victor Hensen Costa Rica expedition (1993/1994). Rev. Biol. Trop. 44 (Suppl. 3): 97-102.         [ Links ]

180. Vargas-Montero, M., Á. Morales-Ramírez & J. Cortés. 2012. Primer informe del  género Gambierdiscus (Dinophyceae)  en el Parque Nacional Isla del Coco, Pacífico Tropical Oriental. Rev. Biol. Trop. 60 (Suppl. 3): 187-199.         [ Links ]

181. Víquez,  R. & P.E. Hargraves. 2009.  Phytoplankton. Text: 97-108, Species list, CD: 1-16. In: I.S. Wehrtmann  &  J.  Cortés  (Eds.).  Marine  Biodiversity  of Costa Rica, Central America. Springer, Berlin.         [ Links ]

182. Waller, T.R. 2007. The evolutionary and biogeographic origins of the endemic  Pectinidae (Mollusca: Bivalvia) of the Galapagos Islands. J. Paleontol. 81: 929-950.         [ Links ]

Wehrtmann, I.S. & J. Cortés (Editors). 2009. Marine Biodiversity of Costa Rica, Central America. Monographiae Biologicae, Volumen 86. Springer + Business Media B. V.,  Berlin. Text: 538  p., List of species in Compact Disk: 500 p.         [ Links ]

Wehrtmann, I.S., J. Cortés & S. Echeverría-Sáenz. 2009. Marine biodiversity of Costa Rica: perspectives and conclusions, Chapter V, 521-533. In: I.S. Wehrtmann and J.  Cortés  (Eds.). Marine Biodiversity of  Costa Rica, Central America. Springer, Berlin.         [ Links ]

183. Wicksten, M.K. & R. Vargas. 2001. A new species of Thor Kingsley, 1878 (Crustacea: Decapoda: Caridea: Hippolytidae) from the tropical eastern Pacific. Proc. Biol. Soc. Washington 114: 139-144.         [ Links ]

184. Williams, A.B. 1986. Mud shrimps, Upogebia, from the  eastern  Pacific  (Thalassinoidea:  Upogebiidae). San Diego Soc. Nat. Hist. Mem. 14: 1-60.         [ Links ]

185. Williamson, S.J., D.B. Rusch, S. Yooseph, A.L. Halpern, K.B. Heidelberg, J.I. Glass, C. Andrews-Pfannkoch, D. Fadrosh, C.S. Miller, G. Sutton, M. Frazier, J.C. Venter. 2008. The Sorcerer II Global Ocean Sampling Expedition:  Metagenomic characterization of viruses within aquatic microbial samples. PLoS ONE 3(1): e1456. doi:10.1371/journal.pone.0001456.         [ Links ]

186. Wilson, H. V. 1904. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the US Fish Commission Steamer Albatross, during 1891, Lieut.  Commander Z.L. Tanner U.S.N.,  commanding. XXX.  The  sponges.  Mem. Mus. Comp. Zoöl., Harvard Coll. XXX: 1-164.         [ Links ]

187. Young, B.E., K. Easley, R. Garrigues, B. Mactavish, P. Murgatroyd & J.R. Zook.  2010. Swallow-tailed gull Creagrus  furcatus in Costa Rica. Cotinga 32: 24-26.         [ Links ]

188. Zullo, V.A. 1991. Zoogeography of  shallow-water cirriped fauna of the Galápagos Islands and adjacent regions in the Tropical Eastern Pacific. Pp. 173-192. In:  M.J.  James  (Ed.).  Galápagos  Marine  Invertebrates: Taxonomy,  Biogeography, and Evolution in Darwin’s  Islands.  Plenum  Press,  New  York  and London.         [ Links ]

*Correspondencia a:
Jorge Cortés. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Ciudad de la Investigación, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica; jorge.cortes@ucr.ac.cr. Escuela de Biología, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica

1. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Ciudad de la Investigación, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica; jorge.cortes@ucr.ac.cr
2. Escuela de Biología, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica
Note: Number references as used in the Tables.

Received 05-I-2012. Corrected 01-VIII-2012. Accepted 24-IX-2012

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